Comparative Anatomical Studies on Some Species of the Genus Amaranthus (Family: Amaranthaceae) for the Development of an Identification Guide Q ⇑ Abbas A
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Annals of Agricultural Science xxx (2017) xxx–xxx Contents lists available at ScienceDirect Annals of Agricultural Science journal homepage: Comparative anatomical studies on some species of the genus Amaranthus (Family: Amaranthaceae) for the development of an identification guide q ⇑ Abbas A. El-Ghamery a, Ahmed M. Sadek a, , Ola H. Abd El Bar b a Botany and Microbiology Department, Faculty of Science, Al-Azhar University, Cairo, Egypt b Agriculturale Botany Department, Faculty of Agriculture, Ain-Shams University, Cairo, Egypt article info abstract Article history: A study of anatomical features of mature leaves and stems (at fruiting stage) of 12 Amaranthus taxa Received 20 September 2016 (Family: Amaranthaceae) shows high variation between them and supplied new characters. The internal Received in revised form 2 November 2016 structures were evaluated to clarify their effectiveness in solving taxonomic complexity and identifica- Accepted 16 November 2016 tion difficulty in this genus. Observation of the transections of blades showed that the epidermis is unis- Available online xxxx eriate, ground tissue consists of angular collenchyma and thin parenchyma. The vascular bundles shape has three patterns crescent, ring, ovate. Also they may be united or separated while the midrib shape in Keywords: cross section has two patterns in which U-shaped, cordate or crescent bundle occurs. All leaves are peti- Amaranthus olate. The examination of the petioles exhibits new and varied characters such as petiole shape (cross sec- Leaf and stem anatomy DELTA key tion), vascular bundles (shape, number, arrangement). While the resulted characters from the Identification observation of the stem structure showed less variation. Nineteen qualitative characters with 38 charac- ter states resulted from leaf anatomy. Only (8) characters were sufficient to generate an identification anatomical key. DELTA program was used in key-generation. Also different measurements were carried out by a photo analysis program (Image J), such as lamina thickness, mesophyll thickness, area of upper and lower epidermal cells and thickness of upper and lower epidermal cells to exhibit most possible dis- similarities between the studied species. Ó 2017 Production and hosting by Elsevier B.V. on behalf of Faculty of Agriculture, Ain Shams University. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc- nd/4.0/). Introduction misapplication of names create intricate problems in its taxonomy (Costea et al., 2001; Iamonico, 2014a, 2014b, 2016a, 2016b; Nestor, The number of Amaranthus species varies in literature. The 2015). genus Amaranthus L. (Amaranthaceae Juss.) consists about 65–80 Furthermore, some Amaranthus species are aggressive annual (Bojian et al., 2003; Mosyakin and Robertson, 1996; Mujica and weeds in the world, causing impacts on the natural floras and veg- Jacobsen, 2003; Iamonico, 2014a, 2014b.). 455 species names for etation and on cultivated field Amaranthus presence can lower the genus Amaranthus are known with the following statistics: grain yield and quality as well as hinder mechanical harvest 105 (23.1%) accepted, 262 (57.6%) synonyms and 88 (19.3%) unas- (Wax, 1995). Some of this species also have been shown to possess sessed (The plant list, 2013). However due to the few studies on allelopathic chemicals that reduce seedling vigor of several crops Amaranthus systematics the number still tentative. Also the and weed species (Menges, 1987, 1988), while others may induce nomenclature of Amaranthus is critical since hundred names were toxicosis and death in dairy cattle (Kerr and Keleh, 1998). The grain published during the centuries and the nomenclatural disorder/ Amaranths are ancient crops with increasing prospects as potential food and feed resources because of their high grain protein and starch quality and high-nitrogen, highly digestible vegetative tis- Peer review under responsibility of Faculty of Agriculture, Ain-Shams University. ⇑ sues (Cai et al., 1998a,b). Some Amaranthus species have medicinal Corresponding author. importance as A. spinosus (Pal et al., 2013). E-mail addresses: [email protected], [email protected] (O.H. Abd El Bar). http://dx.doi.org/10.1016/j.aoas.2016.11.001 0570-1783/Ó 2017 Production and hosting by Elsevier B.V. on behalf of Faculty of Agriculture, Ain Shams University. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/). Please cite this article in press as: El-Ghamery, A.A., et al. Comparative anatomical studies on some species of the genus Amaranthus (Family: Amaran- thaceae) for the development of an identification guide. Ann. Agric. Sci. (2017), http://dx.doi.org/10.1016/j.aoas.2016.11.001 2 A.A. El-Ghamery et al. / Annals of Agricultural Science xxx (2017) xxx–xxx Among the morphological studies on different Amaranthus spe- were immediately fixed in (FAA) according to Johasen (1940). cies; Abdul Rahaman and Oladele (2003) and Gaafar et al. (2015) The third node and internode were selected for the stem study. studied the stomata; Alege and Daudu (2014), Pal et al. (2013), Serial transverse sections (10–20 lm in thickness) were cut with Munir et al. (2011), Hong et al. (2005) studied the leaves. a steel blade on an IHC World KD-1508A rotary microtome and Kraehmer (2013), Carlquist (2003) and Metcalfe & Chalk (1950, fixed on slides by means of Haupt’s adhesive. The sections were 1979) stained with a safranin-fastgreen combination, and then mounted As a consequence, some issues of the taxonomy of Amaranthus in Canada balsam (Sass, 1961). still remain not completely solved, although some approaches Observation and photomicrographs were achieved using XSZ- was carried out (Hauptli and Jain, 1978; Joshi and Rana, 1991; N107 Research Microscope fitted with Premiere MA88-900 digital Bojian et al., 2003, Mosyakin and Robertson, 2003; Hussein, camera. 14 measurements for leaf and stem internal structures, 2005; Pinto and Velasquez, 2010; Iamonico, 2012; Iamonico and epidermal cell area were carried out by the ImageJ Ver. 1.49o pro- Das, 2014). gram (standard software in scientific image analysis) and cali- Because of the poor published data about the leaf and stem ana- brated by using a standard stage micrometer slide. Each recorded tomies, especially concerning the taxonomical implications, we measurement is the average of ten measurements. here present a study aims to evaluate the taxonomic value of leaf Also stomatal density and stomatal index where calculated as and stem structures of some Amaranthus species, with the final described by Wilkinson (1979). aim to facilitate identification Amaranthus and improve the taxon- omy of the genus. SI ¼ S Â 100=S þ E Various data obtained from the description of the leaf anatomy Material and methods were subjected to automated key generation using version 4.12 of the DELTA suite of program (Dallwitz et al., 2000). Seeds of 11 Amaranthus accessions were supplied by the Royal Botanic Garden at Kew (London, UK) while one taxon (A. tricolor) was selected from one of the Botanical Gardens in Egypt Results (National Garden in Nasr City). 12 species of Amaranthus were studied in the present study (Table 1). All the plants were grown The observation of the internal structure of the leaf and stem of from seeds. the studied species revealed (14) qualitative characters (Table 3). The leaf structure was examined using five segments (lengths of 28 qualitative characters were resulted and summarized in 1–2 mm) of the blade and petiole (fruiting stage) and then they (Table 2). Also they comprehensively listed in the following text: Table 1 Species and accessions of Amaranthus used in this study. No. Species Accession no. Origin Subgenus Habitat 1 A. albus L. 0022453 Germany Albersia Wild 2 A. blitum L. 0006563 France Albersia Cultivated 3 A. caudatus L. 0003919 Unknown Amaranthus Wild 4 A. deflexus L. 0020275 Italy Albersia Cultivated 5 A. dubius Mart.ex Thell. 0006574 Colombia Albersia Wild 6 A. graecizans L. 0020242 Portugal Amaranthus Wild 7 A. hybridus L. 0019657 Bolivia Amaranthus Wild 8 A. powellii S. Watson. 0030999 France Amaranthus Wild 9 A. retroflexus L. 0008394 Italy Amaranthus Wild 10 A. spinosus L. 0017882 Unknown Amaranthus Wild 11 A. tricolor L. Unknown Egypt Amaranthus Cultivated 12 A. x ozanonii Piszter. 0008383 Unknown Amaranthus Cultivated Table 2 Qualitative characters of leaf and stem of the studied species of Amaranthus. Character number is preceded by # and states of the same character are assigned serial numbers. Characters Species A. A. A. A. A. A. A. A. A. A. A. A. x albus blitum caudatus deflexus dubius graecizans hybridus powellii retroflexus spinosus tricolor ozanonii #1. Petiole shape in TS/1.crescent/2. 11 4 1 1 1 1 1 1 2 2 3 rounded/3.cordate/4.V-shaped #2. Protuberances/1.present/2.absent 1 1 1 1 1 1 1 1 1 1 2 1 #3. Epidermis type/1.uniseriat with 11 1 1 1 1 1 1 1 1 1 1 radial or pueblos cells #4. Ground tissue type/ 11 1 1 1 1 1 1 1 1 1 1 1.parenchymatous and angular collenchyma #5. Sand crystals in petiole/1.present/2. 11 1 1 1 1 1 1 1 1 1 1 absente #6. Number of vascular bundles/1.8/ 1 2 3 4 3 5 4 6 1 7 6 7 Please cite this article in press as: El-Ghamery, A.A., et al. Comparative anatomical studies on some species of the genus Amaranthus (Family: Amaran- thaceae) for the development of an identification guide. Ann. Agric. Sci. (2017), http://dx.doi.org/10.1016/j.aoas.2016.11.001 A.A. El-Ghamery et al. / Annals of Agricultural Science xxx (2017) xxx–xxx 3 Table 2 (continued) Characters Species A. A. A. A. A. A. A. A. A. A. A. A. x albus blitum caudatus deflexus dubius graecizans hybridus powellii retroflexus spinosus tricolor ozanonii 2.4/3.13/4.6/5.12/6.11/7.7 #7.